(CANCER RESEARCH 52. 3409-3417. June 15, 1992]
Resistance to /V-Benzyladriamycin-14-valerate in Mouse J774.2 Cells:
P-Glycoprotein Expression without Reduced jV-Benzyladriamycin-
14-va lenite Accumulation1
Leonard Lothstein,2 Trevor W. Sweatman, Michael E. Dockter, and Mervyn Israel
Departments of Pharmacology [L. L., T. W. S., M. I.] and Medicine [M. E. D.J, College of Medicine, and Cancer Center [L. L., T. W. S., M. 1.], University of Tennessee-
Memphis, Memphis, Tennessee 38163
ABSTRACT
Ar-Benzyladriamycin-14-valerate (AD 198) ¡s a highly lipophilic ana
logue of Adriamycin with novel cytotoxic mechanisms, greater in vivo
antitumor activity, and the ability to circumvent multidrug resistance due
to P-glycoprotein-mediated drug efflux or decreased topoisomerase II
activity. To identify the mechanism(s) which may confer AD 198 resist
ance, J774.2 mouse macrophage-like cells were selected for growth in
cytotoxic levels of AD 198 (AD 198"). AD 198" cells exhibited over-
expression of the mtlrlb (P-glycoprotein) gene, cross-resistance to Adri-
amycin and vinblastine, and potentiation of drug cytotoxicity by verapa-
mil. However, net intracellular accumulation of AD 198 in AD 198" cells
was unchanged compared to parental cells, while Adriamycin and vin
blastine accumulations were reduced 40% and 95%, respectively. AD 198
was localized in the perinuclear region of the cytoplasm in both parental
and AD 198" cells, with additional vesicular compartmentalization in AD
198R cells. Verapamil-induced reversal of AD 198 resistance coincided
with some drug redistribution from cytoplasmic vesicles, but without
redistribution of AD 198 into the nucleus. These results suggest that AD
198 resistance was not conferred through a P-glycoprotein-mediated
reduction in intracellular drug accumulation but through other cyto
plasmic mechanisms, including, but not limited to, drug compartmental
ization.
INTRODUCTION
The anthracycline antibiotic ADR1 is an effective cancer
chemotherapeutic agent that has been widely used against a
variety of human malignancies (1). Concomitant with its anti-
tumor activity, ADR can produce adverse systemic effects,
including acute myelosuppression, cumulative cardiotoxicity,
and gastrointestinal toxicity (2). At the cellular level, in both
cultured mammalian cells and primary tumor cells, ADR can
select for multiple mechanisms of drug resistance that decrease
its chemotherapeutic efficacy. These mechanisms include P-gp-
mediated MDR, characterized by the energy-dependent trans
port of drugs from the cell (3), and multidrug resistance con
ferred by decreased topoisomerase II activity, resulting in de
creased ADR-induced DNA strand scission (4-6). Among the
potential avenues of circumvention of systemic toxicity and
cellular drug resistance is the development of semisynthetic
Received 3/11/91; accepted 4/3/92.
The costs of publication of this article were defrayed in part by the payment
of page charges. This article must therefore be hereby marked advertisement in
accordance with 18 U.S.C. Section 1734 solely to indicate this fact.
1Supported in part by American Cancer Society Grants IN-176-A and JFRA-
287 (L. L.), Pharmaceutical Manufacturers Association Foundation Research
Starter Grant (L. L.), and NIH-National Cancer Institute Research Grants
CA44890 (T. W. S.), CA37082 (M. I.), and CA37209 (M. I.).
2To whom requests for reprints should be addressed, at Department of
Pharmacology, University of Tennessee College of Medicine, 874 Union Avenue,
Memphis, TN 38163.
3The abbreviations used are: ADR, Adriamycin hydrochloride (doxorubicin);
AD 198, /V-benzyladriamycin-14-valerate; AD 288, A'-benzyladriamycin; DMSO,
dimethyl sulfoxide; MDR, multidrug resistance (resistant); P-gp, P-glycoprotein;
VBL, vinblastine sulfate; VRP, (±)verapamil; cDNA, complementary DNA;
HPLC, high-performance liquid chromatography; IC50,drug concentration inhib
iting growth by 50% after 72 h of continuous exposure; TBS, 20 mm Tris-HCl,
pH 7.5, 0.5 M NaCl; TTBS, 0.2% Tween 20 in TBS; PBS, phosphate-buffered
saline.
ADR analogues which demonstrate greater tumor-specific tox
icity and less susceptibility to various forms of resistance. One
such highly hydrophobic analogue, AD 198, exhibits a variety
of mechanistic differences compared with ADR, including
weaker binding to purified DNA, preferential inhibition of
RNA versus DNA synthesis, irreversible G2/M blockade, pro
nounced membrane lytic activity, and a lack of inhibition of
purified mammalian topoisomerase II despite significant levels
of protein-associated DNA strand breaks in alkaline elution
assays (2, 7-9). When compared with ADR, AD 198 demon
strates enhanced cytotoxicity against cultured murine and hu
man tumor cells and the ability to circumvent MDR in P388
and LI210 leukemic cells and B16-BL6 melanoma cells, and
both MDR and resistance due to altered topoisomerase II
activity in variant CCRF-CEM leukemic cells (10, 11). Despite
the high degree of toxicity seen in vitro, AD 198 exhibits limited
efficacy against transplanted MDR LI210 cells in vivo. This
observation suggests that resistance to AD 198 may be con
ferred either systemically through enhanced drug metabolism
or pharmacologie sanctuary of the neoplasia or through cellular
resistance (10).
In an effort to identify the mechanism(s) by which mamma
lian cells may acquire resistance to AD 198, the selection of
resistant variants of the mouse macrophage-like cell line J774.2
were attempted by continuous exposure to increasing concen
trations of AD 198. J774.2 cells are amenable to the rapid
selection of drug-resistant variants, a number of which have
been extensively analyzed (12-17). Selection of variant J774.2
cells would represent the first reported selection of myeloid
cells for resistance to AD 198. In this report, it is shown that
(a) AD 198-resistant variants of J774.2 (AD 198R) can be
rapidly selected; (b) AD 198Rcells exhibit several characteristics
of MDR, including P-gp overexpression, cross-resistance to
other drugs, and attenuation of resistance by the calcium chan
nel blocker VRP; and (c) unlike cross-resistance to ADR and
VBL, AD 198 resistance is not associated with reduced net AD
198 accumulation.
MATERIALS AND METHODS
Materials. AD 198 (Fig. 1), [14C]AD 198 (3.8 mCi/mmol), and [I4C]
ADR (5.05 mCi/mmol) were prepared as described previously (8, 11).
VBL and VRP were obtained from Sigma Chemical Co. (St. Louis,
MO). ADR was a generous gift from Farmitalia Carlo Erba (Milan,
Italy). [3H]VBL (11.2 Ci/mmol) was obtained from Amersham Corp.
(Arlington Heights, IL), and [-"P]dCTP (3000 Ci/mmol) was from
NEN/DuPont (Boston, MA). Drug-sensitive J774.2 cells, VBL-resist-
ant J774.2 cells (13, 17), and mdr cDNA probes (14) were generously
supplied by Dr. Susan Honvitz (Albert Einstein College of Medicine,
Bronx, NY).
Cell Culture. The propagation J774.2 cells and the selection of VBL-
resistant J7.V3-0.04 cells have been described previously (13, 14, 18).
AD 198R variants were selected by stepwise, continuous exposure of
J774.2 cells to increasing concentrations of AD 198, beginning with 50
3409
Research.
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